Many of the bolded characters in the characterization above are apomorphies of subsets of streptophytes along the lineage leading to the embryophytes, not apomorphies of crown-group embryophytes per se.

All groups below are crown groups, nearly all are extant. Characters mentioned are those of the immediate common ancestor of the group, [] contains explanatory material, () features common in clade, exact status unclear.

Age. K. Bremer et al. (2004) suggested an age of about 111 m.y. for this node, Magallón et al. (2015) an age of ca 81.8 m.y., and Wikström et al. (2015) an age of (104-)92(-79) m.y.; around 94.2 m.y. is the estimate in Tank et al. (2015: Table S2) and (101.4-)86.6(-71.8) m.y. in Tank and Olmstead (2017).

Age. K. Bremer et al. (2004) suggested an age of about 101 m.y. for crown-group Paracryphiaceae/Paracryphiales; for more details, see below.

Note:Boldface denotes possible apomorphies, (....) denotes a feature common in the clade, exact status uncertain, [....] includes explanatory material. Note that the particular node to which many characters, particularly the more cryptic ones, should be assigned
is unclear. This is partly because homoplasy is very common, in addition, basic information for all too many characters is very incomplete, frequently coming from taxa well embedded in the clade of interest and so making the position of any putative apomorphy uncertain. Then there are the not-so-trivial issues of how character states are delimited and ancestral states are reconstructed (see above).

Evolution:Divergence & Distribution. The three genera included in Paracryphiales are quite different florally and are also rather unlike other campanulids. Quintinia is reported to have ellagic acid (along with iridoids!), and it and fossils that have been associated with it may even have bitegmic ovules (Friis & Pedersen 2012). Sphenostemon and Paracryphia sometimes have more than twice as many stamens as perianth parts; the apomorphies of Paracryphia in particular represent a very odd combination for an euasterid. Altogether, something of a conundrum.

Chemistry, Morphology, etc. For more on the inferior/superior ovary distinction in the campanulids, see the Asterales page.

All three genera are poorly known - Sphenostemon particularly so - and need an exhaustive comparative study; this should be accompanied by a reworking of their molecular relationships.

Phylogeny. For the relationships of Paracryphiales, see the asterid II clade.

3 [list]/36 - as genera below. Southwest Pacific: Philippines to New Zealand and New Caledonia.

Age. K. Bremer et al. (2004) suggested an age of about 101 m.y. and Wikström et al. (2015) an age of (98-)78(-32) m.y. for crown-group Paracryphiaceae.

The Late Cretaceous Silvianthemum suecicum, from rocks in southern Sweden has several exquisitely-preserved features - perhaps most notably the radial stylar cells - suggesting a relationship with Quintinia in particular (Friis 1990). Martínez-Millán (2010: 83.5 m.y.o.) recovered this relationship in a phylogenetic analysis of possibly related campanulids, although no other Paracryphiales were included, and she thought better knowledge of Escalloniales might help place the fossil more securely (see also Friis et al. 2011, 2013b). However, Silvianthemum has tricolp(or)ate pollen, there are eight stamens (c.f. Friis et al. 2011: the perianth is 5-merous), the anthers appear to be dorsifixed (a synapomorphy with Quintinia - Martínez-Millán 2010), there are three short, adaxially grooved styles (c.f. Quintinia), and three stipitate parietal placentae. Bertilanthus scanicus, from the same rocks, has a very similar set of characters, and, like Quintinia, it has glandular hairs, but it also has stamens opposite the petals, a feature at most vanishingly infrequent in the campanulids, and differs in other respects from Quintinia (Friis & Pedersen 2012). However, the androecium in extant Paracryphiales is variable. The ovules of Bertilanthus may be bitegmic ("ovary wall [sic] split in two": Friis & Pedersen 2012: p. 326), which would be in line with reports for Quintinia.

If the relationships of these fossils hold, then the current distribution of Paracryphiales has little to do with their past distribution. However, a position of the fossils within Cornales was suggested by Beaulieu et al. (2013a), which agrees better with their morphology (and geography). Moreover, Friis et al. (2013b) are inclined to question the position of Quintinia itself in the campanulids.

Quintinia is an evergreen tree or shrub with spiral, serrate, estipulate leaves; the plant is covered with peltate scales. The inflorescence is terminal and the flowers have free sepals and petals and an inferior ovary. The fruit is a septicidal capsule.

Sphenostemon is usually a small, evergreen tree with spiral, strongly serrate leaves that lack much in the way of stipules, and terminal inflorescences with rather small flowers. The sepals and petals are free, the anther thecae are borne on stout filaments, and there is no style; the fruit is fleshy and two-seeded.

Evolution:Divergence & Distribution. The vessel elements of the three genera are very long and the perforation plates have many bars, indeed, aspects of the wood anatomy of Paracryphia have been considered to be among the most primitive in angiosperms.

Chemistry, Morphology, etc. The considerable variation in petiole anatomy in Sphenostemon may depend where in the petiole the transverse sections were taken. Styloids are visible on the abaxial surface
of the lamina of Papuasian species of Sphenostemon; they look rather like cystoliths.

The placentation of Quintinia is basically parietal (Bensel & Palser 1975b; see also Friis et al. 2103b); the ovules
may be bitegmic and crassinucellate (Mauritzon 1933; Philipson 1974; Friis et al. 2013b). For some details of the flower of Sphenostemon, see Endress (2008c). Its fruit is often described as being a drupe, but Lundberg (2001c) characterized it as being a pseudo-drupe (and the seeds as being pachychalazal), while Savinov (2003) described it as being a berry. The nature of the perianth in both Paracryphia and Sphenostemon needs attention. In the former, it is almost as if the bract encloses the sessile flower, while in the latter the first perianth members are lateral and subpeltate.

For Paracryphia, see also
Dickison and Baas (1977) and Carlquist (2012c) for wood anatomy, and Swamy (1953b), Lundberg (2001e) and Dickison and Lunberg (2016), all general. For Quintinia, see Lundberg (2000d) and Dickison and Lunberg (2016), both general. For Sphenostemon, see Bailey and Swamy (1953), Jéremie (1997) and Thiv (2016) for general
information, Carlquist (2012c) for wood anatomy, and Savinov (2003) for fruit and seed anatomy.

Classification. The three genera have all been placed in monogeneric families, but only relatively recently; they are combined here (see also A.P.G. 2009, 2016).

Thanks. Thanks to Y. Pillon for comments.

Previous Relationships. Just about everywhere. Paracryphiaceae were included in Theales
by Cronquist (1981) and in Theanae by Takhtajan (1997). Quintinia has long been included in woody Saxifragaceae/Hydrangeaceae. Baas (1975) thought that
Sphenostomonaceae (and Phellinaceae) were members of Celastrales, close to
Icacinaceae; Cronquist (1981) had placed them in Aquifoliaceae, next to Icacinaceae, while Takhtajan (1997) included them in Icacinales. Sphenostemon (as Idenburgia) has also been placed in Trimeniaceae.